What Is Evolution?
Are Humans Evolving?
Essays & Articles
Central Dogma of Molecular Biology
How Many Genes Do We Have?
Random Genetic Drift
Evolution and Abiogenesis
Random Genetic Drift
v1.3 ©2006 Laurence A. Moran
Science v. Religion
Another curious aspect of the theory of evolution
is that everybody thinks he understands it. I mean philosophers, social
scientists, and so on. While in fact very few people understand it, actually,
as it stands, even as it stood when Darwin expressed it, and even less as we
now may be able to understand it in biology.
he two most important mechanisms of evolution are natural selection and random genetic drift. While most people have a reasonable understanding of natural selection, they don't realize that random genetic drift is also important. The anti-evolutionists, in particular, tend to focus their attacks on natural selection, not realizing that there is much more to modern evolutionary biology.
Charles Darwin didn't know about random genetic drift but we now recognize drift as an important feature of evolution. This is one of the reasons why most modern evolutionary biologists no longer think of themselves as strict "Darwinists". When you hear the anti-evolutionists equating "evolution" with "Darwinism" you know they haven't done their homework! (It's very unusual to find an anti-evolutionist who understands random genetic drift or any of the other modern concepts in evolution. As a matter of fact, it's rare to find an anti-evolutionist who even understands the 145-year old concept of natural selection.)
Random genetic drift is a stochastic process (by definition). A classic example of random genetic drift is the random nature of transmitting alleles from one generation to the next given that only a fraction of all possible zygotes become mature adults. The easiest case to visualize is the one that involves binomial sampling error. If a pair of diploid sexually reproducing parents (such as humans) have only a small number of offspring then not all of the parent's alleles will be passed on to their progeny due to chance assortment of chromosomes at meiosis. In a large population this will not have much effect in each generation because the random nature of the process will tend to average out. But in a small population the effect could be rapid and significant.
Suzuki et al. explain it as well as anyone I've seen;
If a population is finite in size (as all populations are) and if a given pair of parents have only a small number of offspring, then even in the absence of all selective forces, the frequency of a gene will not be exactly reproduced in the next generation because of sampling error. If in a population of 1000 individuals the frequency of "a" is 0.5 in one generation, then it may by chance be 0.493 or 0.505 in the next generation because of the chance production of a few more or less progeny of each genotype. In the second generation, there is another sampling error based on the new gene frequency, so the frequency of "a" may go from 0.505 to 0.501 or back to 0.498. This process of random fluctuation continues generation after generation, with no force pushing the frequency back to its initial state because the population has no "genetic memory" of its state many generations ago. Each generation is an independent event. The final result of this random change in allele frequency is that the population eventually drifts to p=1 or p=0. After this point, no further change is possible; the population has become homozygous. A different population, isolated from the first, also undergoes this random genetic drift, but it may become homozygous for allele "A", whereas the first population has become homozygous for allele "a". As time goes on, isolated populations diverge from each other, each losing heterozygosity. The variation originally present within populations now appears as variation between populations.
Suzuki, D.T., Griffiths, A.J.F., Miller, J.H. and
Random genetic drift is not limited to species that have few offspring, such as humans. In the case of flowering plants, for example, the stochastic element is the probabilty of a given seed falling on fertile ground. In the case of some fish and frogs it is the result of chance events that determine whether a newly hatched individual will survive. Drift is also not confined to diploid genetics; it can explain why we all have mitochondria that are descended from those of a single women who lived hundreds of thousands of years ago.
This does not mean that there was a single female from whom we are all descended, but rather that out of a population numbering perhaps several thousand, by chance, only one set of mitochondrial genes was passed on. (This finding, perhaps the most surprising to us, is the least disputed by population geneticists and others familiar with genetic drift and other manifestations of the laws of probability.)
Curtis, H. and Barnes, N.S. in Biology 5th ed. Worth Publishers (1989 p.1050)
But random genetic drift is even more than this. It also refers to accidental random events that influence allele frequency. For example,
Chance events can cause the frequencies of alleles in a small population to drift randomly from generation to generation. For example, consider what would happen if [a]... wildflower population ... consisted of only 25 plants. Assume that 16 of the plants have the genotype AA for flower color, 8 are Aa, and only 1 is aa. Now imagine that three of the plants are accidently destroyed by a rock slide before they have a chance to reproduce. By chance, all three plants lost from the population could be AA individuals. The event would alter the relative frequency of the two alleles for flower color in subsequent generations. This is a case of microevolution caused by genetic drift...
Disasters such as earthquakes, floods, or fires may reduce the size of a population drastically, killing victims unselectively. The result is that the small surviving population is unlikely to be representative of the original population in its genetic makeup - a situation known as the bottleneck effect.... Genetic drift caused by bottlenecking may have been important in the early evolution of human populations when calamities decimated tribes. The gene pool of each surviving population may have been, just by chance, quite different from that of the larger population that predated the catastrophe.
Campbell, N.A. in Biology 2nd ed. Benjamin/Cummings (1990 p.443)
Several examples of bottlenecks have been inferred from genetic data. For example, there is very little genetic variation in the cheetah population. This is consistant with a reduction in the size of the population to only a few individuals - an event that probably occurred several thousand years ago. An observed example is the northern elephant seal which was hunted almost to extinction. By 1890 there were fewer than 20 animals but the population now numbers more than 30,000. As predicted there is very little genetic variation in the elephant seal population. It is likely that the twenty animals that survived the slaughter were more "lucky" than "fit".
Another example of genetic drift is known as the founder effect. In this case a small group breaks off from a larger population and forms a new population. This effect is well known in human populations;
The founder effect is probably responsible for the virtually complete lack of blood group B in American Indians, whose ancestors arrived in very small numbers across the Bering Strait during the end of the last Ice Age, about 10,000 years ago. More recent examples are seen in religious isolates like the Dunkers and Old Order Amish of North America. These sects were founded by small numbers of migrants from their much larger congregations in central Europe. They have since remained nearly completely closed to immigration from the surrounding American population. As a result, their blood group gene frequencies are quite different from those in the surrounding populations, both in Europe and in North America.
The process of genetic drift should sound familiar. It is, in fact, another way of looking at the inbreeding effect in small populations ... Whether regarded as inbreeding or as random sampling of genes, the effect is the same. Populations do not exactly reproduce their genetic constitutions; there is a random component of gene-frequency change.
Suzuki et al. op. cit.
There are many well studied examples of the founder effect. All of the cattle on iceland, for example, are descended from a small group that were brought to the island more than one thousand years ago. The genetic make-up of the icelandic cattle is now different from that of their cousins in Norway but the differences agree well with those predicted by genetic drift. Similarly, there are many pacific islands that have been colonized by small numbers of fruit flies (perhaps one female) and the genetics of these populations is consistant with drift models.
These examples, and many more, show that it is wrong to consider natural selection as the ONLY mechanism of evolution and it is also wrong to claim that natural selection is the predominant mechanism. This point is made in many genetics and evolution textbooks. For example;
One of the most important and controversial issues in population genetics is concerned with the relative importance of genetic drift and natural selection in determining evolutionary change. The key question at stake is whether the immense genetic variety which is observable in populations of all species is inconsequential to survival and reproduction (ie. is neutral), in which case drift will be the main determinant, or whether most gene substitutions do affect fitness, in which case natural selection is the main driving force. The arguments over this issue have been intense during the past half- century and are little nearer resolution though some would say that the drift case has become progressively stronger. Drift by its very nature cannot be positively demonstrated. To do this it would be necessary to show that selection has definitely NOT operated, which is impossible. Much indirect evidence has been obtained, however, which purports to favour the drift position. Firstly, and in many ways most persuasively is the molecular and biochemical evidence...
Harrison, G.A., Tanner, J.M., Pilbeam, D.R. and Baker, P.T.
The book by Harrison et al. is quite interesting because it goes on for several pages discussing the controversy. The authors point out that it is very difficult to find clear evidence of selection in humans (the sickle cell allele is a notable exception). In fact, it is difficult to find good evidence for selection in most organisms. (All of the arguments are "after the fact," but probably correct!)
One of the consequence of fixation by random genetic drift is that beneficial alleles can be lost by drift before they are fixed by natural selection. Take the example of an allele with a 5% selective advantage. The probability of fixation of this allele is only 10% and not 100% as most people imagine. The difference is due to the same stochastic events that govern random genetic drift. In 90% of the cases, the less beneficial allele will reach fixation. Similarly, new detrimental alleles are not always eliminated from the population ....
In any population, some proportion of loci are fixed at a selectively unfavorable allele because the intensity of selection is insufficient to overcome the random drift to fixation. Very great skepticism should be maintained toward naive theories about evolution that assume that populations always or nearly always reach an optimal constitution under selection. The existence of multiple adaptive peaks and the random fixation of less fit alleles are integral features of the evolutionary process. Natural selection cannot be relied on to produce the best of all possible worlds.
Suzuki, D.T., Griffiths, A.J.F.,
Miller, J.H. and Lewontin, R.C.
The relative importance of drift and selection depends, in part, on estimated population sizes. For any given set of alleles, random genetic drift is much more important in small populations than in large populations. It is important to remember that most species consist of numerous smaller inbreeding populations called "demes". It is these demes that evolve and that makes drift very important in the real world.
The effect of population size is not important when you only consider overall change and not the effect on a single allele. This is because larger populations have more mutations and the probability of fixation by random genetic drift is identical to the rate of mutation for most alleles. Thus, large and small populations evolve at the same rate by random genetic drift.
Studies of evolution at the molecular level have provided strong support for drift as a major mechanism of evolution. Observed mutations at the level of gene are mostly neutral and not subject to selection. One of the major controversies in evolutionary biology is the neutralist-selectionist debate over the importance of neutral mutations. Since the only way for neutral mutations to become fixed in a population is through random genetic drift, this controversy is actually over the relative importance of drift and natural selection.